Journal of Plankton Research Vol.20 no.S pp.

805-816, 1998

Food regulation of growth and maturation in a natural population

of Aurelia aurita (L.)

Haruto Ishii1 and Ulf Bamstedt

Department of Fisheries and Marine Biology, University of Bergen, Bergen High
Technology Center, N-5020 Bergen, Norway

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'Present address and for correspondence: Tokyo University of Fisheries, 4-5-7
Kounan, Minato-ku, Tokyo, 108, Japan

Abstract. Growth and maturity development of the moon jellyfish, Aurelia aurita, were recorded in
Vagsb0pollen, a small and semi-enclosed bay on the Norwegian west coast, and compared to those
of medusae transferred to excess food and starving conditions, respectively. Mesozooplankton were
extremely scarce in Vagsb0pollen. The abundance and biomass of the medusae in the poll were higher
than those typically found in open waters, reaching a maximum of 22 ind. nr-1 and 710 mg C nr 3 in
June. The average diameter of medusae in the poll increased to 8 cm until the last part of June, with
an instantaneous growth rate between 1.S and 20% day 1 , thereafter retarding somewhat, giving a
negative growth rate of up to 2.6% day-'. Starving medusae showed a negative growth rate of up to
13.4% day~', and all the medusae were dead after 49 days. Well-fed medusae showed a very stable
growth over a 56 day period, diverging from the poll population from early June, and with a growth
rate between 3.8 and 9.8% day-'. Medusae from the poll population began carrying planulae on their
oral arms when at least 5 cm in diameter, whereas not even the largest medusa of 15.6 cm diameter
among those in the well-fed group produced any planulae. For the first time, it is thus explicitly shown
that the size and maturity of A.aurita are externally controlled through food availability. Scarcity of
food reduces the growth rate, but also changes the energy allocation towards reproduction, which thus
occurs at a smaller size than for well-fed medusae. Its plasticity makes it possible for this species to
exploit environments with low advection of food and develop high abundance in such environments,
without losing fecundity.

Introduction
The moon jellyfish, Aurelia aurita, is a cosmopolitan species found in the waters
of Japan (Yasuda, 1971; Ishii et a/., 1995; Omori et a/., 1995), as well as in the
coastal waters of Europe (Moller, 1980; Hernroth and Grondahl, 1985; van der
Veer and Oorthuysen, 1985; Schneider, 1989; Lucas and Williams, 1994; Olesen
et a/., 1994; Schneider and Behrends, 1994; Lucas, 1996) and North America
(Hamner and Jenssen, 1974). The pattern of the population dynamics varies, with
the maximum abundance usually being <1 individual (ind.) nr 3 in open coastal
waters (cf. Moller, 1980). Reports from two enclosed areas have indicated
considerably higher abundance, with up to 25 ind. nr 3 (Lucas, 1996) and 300 ind.
m~3 (Olesen et ai, 1994), but the maximum average diameter only reached 5.4 and
3.7 cm, respectively, in these environments, whereas the maximum average diam-
eter reported for more open areas ranges from 20 to 30 cm (cf. Moller, 1980;
Schneider and Behrends, 1994). These findings indicate that food shortage
restricted growth in the densely populated environments, and it has been
assumed that there is a density-dependent regulation mechanism for A.aurita
populations (Schneider and Behrends, 1994). However, although it is a logical
suggestion to explain the small size as an effect of food shortage, it has not
© Oxford University Press 805
HJshii and U.Bimstedt

hitherto been specifically shown that food regulates the individual growth within
such populations. Furthermore, because reproduction, as reported for more open
environments, occurs when the medusa has reached at least 7-8.5 cm diameter
(Yasuda, 1971; Omori et al., 1995), the consequences of the small size in enclosed
areas may also include a dramatic reduction in reproductive output, if size at
maturity is the same as that in the open environments.
Vagsb0pollen is a semi-enclosed bay with an annual population of A.aurita that
occurs in much higher abundance than in the nearby open water (Berstad et al.,

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1995). This environment is connected to open water through a very shallow
channel of -600 m length, leading to Kviturdvikpollen, which in turn is connected
to a deeper channel of -600 m length, leading to more open water (Figure 1).
Although tidal currents cause some renewal of the water (cf. Dybern, 1967),
exchange of medusae between the inner and outer populations is probably very
limited, due to the length and shallowness of the entrance system. This locality
is, therefore, ideal as a study site for the growth and development of A.aurita.
Furthermore, because the density-dependent growth hypothesis (cf. Schneider
and Behrends, 1994) has not been evaluated experimentally, a simple design
where the natural population is compared with well-fed animals would give us
the opportunity to do so. In this paper, we therefore report the results from an
investigation where the growth and development of medusae in Vagsb0pollen
were compared to those of poll medusae transferred either to conditions of excess
food or to starving conditions.

Method
Aurelia aurita was sampled between 29 April and 16 July 1996 in Vagsb0pollen,
south of Bergen, Norway (Figure 1), and all experiments were performed at the
marine biological station of the University of Bergen, situated close to the study
locality. Vagsb0pollen is a semi-enclosed bay with a maximum length/width of
1300/300 m, and a maximum depth of 12 m (Dybern, 1967). Sampling of A.aurita
was conducted weekly in the deepest part by using a 500 um mesh dip net, with a
101 plastic bag as non-filtering cod end. The net was towed for 5 min in the surface
water at a constant speed. Surface water temperature was measured simul-
taneously. Once a month, zooplankton were sampled with a vertical haul, using a
180 um meshed conical net, and the sample was immediately preserved in 5%
formalin for later microscopic analysis at the laboratory. Collected medusae were
kept in carboys of -301 volume, filled with local surface water during transporta-
tion to the laboratory. After returning to the laboratory, less than 1 h after
sampling, the diameter of each individual was measured to the nearest millimeter
using a Wild M5 dissection microscope (small medusae) or directly by eye, using
a millimeter graded ruler, and the presence of eggs and planula larvae olA.aurita
was recorded. Measurements were made by placing an individual medusa with
exumbrella down in a large Petri dish with sea water, avoiding exposure in the air
as far as possible. The abundance of A.aurita was calculated from the number of
sampled medusae by assuming afilteringefficiency of 100% for the full 5 min tow,
giving the volume filtered as net-opening area times towed distance.
806
 Food regulation ot Aurelia

5'10' 5'1S' 5'20" 5"25'E

60'25'N

L
Bergen

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60"20'

Marine
Biological
Station

K»iturdvikpollen ^

60"15f

Fig. 1. Sampling area ot A.aurita in Vagsbopollen, south of Bergen.

The hydrographic conditions, including salinity, were similar to those for

medusae used by Bamstedt (1990), so the dry weight (DW; mg) of individual
A.aurita medusae was estimated from the equation of Bamstedt (1990):

DW = 0.002 x D2m

where D is the diameter in millimeters. The dry weight was converted to carbon
assuming 4.3% of dry weight (Larson, 1986). The instantaneous growth rate (u;
day 1 ) was calculated on the basis of increments in dry weight:
807
H.Ishii and UBfimstedt

where W^ and W2 are the dry weights, and f, and t2 are the day numbers on two
consecutive analyses.
Aurelia aurita for tank experiments were sampled in Vagsb0poIlen on 22 May
1996 in the same way as in the population study. After returning to the labora-
tory, the collected medusae were transferred to outdoor cylindrical tanks of 2001
volume, filled with continuously aerated sea water. The tanks were placed within

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bigger tanks with flow-through sea water, in order to maintain a constant temper-
ature, and the whole construction was covered with a roof, eliminating direct
sunlight. Seven tanks were used, each of them loaded with 8-13 individuals
randomly selected from the stock of collected medusae. Five of the tanks were
given food in excess in order to saturate feeding of the medusae, and the remain-
ing two tanks were kept without any food. Zooplankton as food were collected
by a 300 um meshed net from Raunefjorden close to the laboratory, and
consisted of -90% in volume of copepodids of Calanus finmarchicus and the
lobate ctenophore Bolinopsis infundibulum. The bell diameter and the presence
of eggs and planula larvae of A.aurita were analyzed weekly in the same way as
for the field medusae, and both the water and the food renewed.
The sex of immature individuals was indistinguishable in the dissection micro-
scope. Medusae were therefore classified as follows: (1) males and unripe females
with no visible gametes in gonads; (2) unripe females, but their oocytes visible in
gonads; (3) ripe females with mature oocytes in gonads, and fertilized eggs and
planula larvae in brood sacs on oral arms; (4) ripe females with fertilized eggs and
planula larvae in brood sacs on oral arms, but no visible oocytes in gonads.

Results
Zooplankton sampled with a 180 um meshed net were extremely scarce in the
monthly samples, with a maximum abundance of 6.2 ind. nr 3 , and consisted
mainly of the two hydromedusae Clytia quadrata and Chemisphaerica.
Aurelia aurita were present in Vagsb0pollen throughout the sampling period
and Figure 2 illustrates its average abundance, biomass and instantaneous growth
rate during the period. The temperature rose gradually from April to June
(8-15°C), climbing up to a maximum on 9 July (15.8°C), and with a slight decrease
to 13.5°C on 16 July (Figure 2). The abundance of medusae was high throughout
the period, with the highest values at the start (16 ind. nr 3 in late April) and in
June (4-22 ind. nr 3 ), and with no significant temporal trend. The biomass was low
in April and May, but steadily rising from -4 mg C m~3 on 29 April to
-20 mg C nr 3 on 13 May. Peak biomass for the period was recorded on 24 June,
at -710 mg C nr 3 , after which the biomass decreased to -100 mg C nr 3 when the
study ended on 16 July. The average instantaneous growth rate varied consider-
ably, but showed a clear decreasing temporal trend, with a peak value on 10 May
of 0.25 day 1 , and close to zero during the summer.
The individual size ranged from 0.2 to 2.9 cm- diameter on 29 April, with the
peak in the size class 1.0-2.0 cm diameter, indicating a recruitment mainly before
808
 Food regulation of Aurelia

20

15
co
3
10
cd
pel

5
m
<v
*~ 0

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i 25
a
Cfl 20
c
15
cu
anc

10
-o
c 5
3

0.2

.5 -0.2

Apr Kay Jun Jul

Fig. 2. Water temperature and abundance, biomass and instantaneous growth rate of A.aurila in
Vagsb0pollen.

29 April (Figure 3). The two smallest size classes gradually disappeared, indicat-
ing that strobilation was not continuous. An approximately normal distribution
was sustained throughout the investigation, confirming the homogeneity of the
population, with no recruitment or mixing with other populations. However,
809
HJshii and U.Bamstedt

29 Apr II Jun
80 N=0. 151. 0 30 H=\Z. 172. 51
60 20
40
20 10
0 LJLl
7 Kay 17 Jun
60 N=0. 138. 0 30 NM8. 34. 50

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40 20
20 10
0 0

13 Kay 24 Jun
30 :Q. 66. 0 60 N=I4. 213. 48

20 40
10 20
0 0
c 22 lay 1 Jul
30 N=23. 91. 57 60 N=8. 74. 48

20
10
0

30 Hay 9 Jul
30 !»=20. 71. 53 30 N=0. 100. 44

20 20
10 10
0 JLi -—-^-s 0

3 Jun 16 Jul
N=I9. 79. 53 40 N=0. 39. 31
30 DStarved
20 1 QField

0 2 10 12 14
10
0
0 2 4 6 8
Mi
10 12 14
lell-fed

Bell diameter (cm)

Fig. 3. Percentage frequency distributions of the bell diameter of A.auriia in Vagsb0pollen and of
starved and well-fed ones cultured in the tank. N is the number of starved individuals, those collected
in the field, and well-fed ones, from left to right, respectively. The culture experiments were started
on 22 May and all starved individuals were dead after 1 July. The data of starved ones on 28 June
were not included.

median and mean size only increased until 24 June, when the largest medusa size
of 11.8 cm was observed, thereafter retarding somewhat through a decreased
frequency of the largest medusae.
810
 Food regulation of Aurelia

The size frequency of A.aurita kept in tanks with excess food was similar to the
field sampled population during the three first sampling occasions, but diverged
thereafter, through a higher increase in large medusae and a continuous gradual
increase in the size distribution throughout the study (Figure 3). The size
frequency of A.aurita kept in starving conditions diverged rapidly from the well-
fed group (Figure 3), and all the medusae were shrunk and dead after 1 July.
The mean bell diameters of medusae from the two experimental groups and
from the field population are displayed in Figure 4. All three groups showed iden-

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tical initial values at the start, 22 May, but starving animals reduced size steadily,
showing an average diameter of 2.3 cm after 6 weeks, and were all dead on 9 July.
The average size of the two other groups began to diverge in June and this differ-
ence was exaggerated after 24 June. Well-fed medusae showed a linearly
increased trend in bell diameter, whereas the field medusae stopped growth
completely. A summary of the average instantaneous growth rate of the three
groups at different periods is displayed in Table I.
The relative proportions of females in different maturity stages are given for
the natural field population (Figure 5A) and the experimental well-fed group
(Figure 5B). In the natural population, sexually immature females dominated
until the end of May, when the population was composed of metephyrae and
small medusae. The percentage of sexually mature females increased during June
and July as gonad development progressed. Females with the oocytes in gonads
were first found on 11 June and the percentage increased to 39.9% on 24 June,
thereafter decreasing gradually. Ripe females with planulae were first observed
on 17 June and this stage increased to 43.6% on 16 July. On 9 July, females with
empty gonads and planulae on the oral arms were first observed, and their
percentage increased to 7.7% on 16 July. The minimum bell diameter of females
with oocytes in gonads and with both eggs and planulae was 5.0 cm.
In the experimental well-fed group, females with the oocytes in gonads were
first identified on 24 June, and the percentage increased to 61.3% on 16 July

14

,2
u

-O

<u *•-» r-... _ Starved

C 2
BJ
ID .
2 0

May Jun Jul

Fig. 4. Mean bell diameter of A.aurita in Vagsb0pollen and of well-fed and starved individuals
cultured in the tank.

811
HJshii and U.Bamstedl

Table L Aurelia aurita: instantaneous growth rate within each period in the field and cultured
individuals in the tank
Period Instantaneous growth rate (day 1 )

Field Tank

Starved Well-fed
22-30 May 0.065 -0.020 0.085
30 May-3 June 0.202 0.002 0.097

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3-11 June 0.023 -0.003 0.098
11-17 June 0.015 -0.054 0.062
17-24 June 0.066 -0.040 0.042
24-28 June _ -0.091 _
28 June-1 July _ -0.134
24 June-1 July -0.014 _ 0.055
1-9 July -0.026 - 0.038
9-16 July 0.011 — 0.052

100 4 A

80

60

40-

20

0
May Jun Jul

100

8(H
y
60-

|
20-

0
May Jun Jul

Fig. S. The percentage ratio of (1) males and unripe females with no visible gametes in gonads, (2)
unripe females, but their oocytes visible in gonads, (3) ripe females with mature oocytes in gonads,
and fertilized eggs and planula larvae in brood sacs on oral arms, (4) ripe females with fertilized eggs
and planula larvae in brood sacs on oral arms, but no visible oocytes in gonads, to all A.aurita, in Vags-
b0pollen (A) or in the tank with well-fed individuals (B).

(Figure 5B). No A.aurita with planula larvae were observed in the study period.
The minimum bell diameter of medusae with oocytes in the gonads was 9.4 cm.
Not even the largest females with a bell diameter of 15.6 cm had developed
812
 Food regulation of Aurelia

planula larvae. In the experimental starved group, no A.aurita with the oocytes
in gonads or planulae were observed.
If we assume that all medusae in category 1 (Figure 5) from 16 July were males,
the proportion of males in the field population was only 15.4% and that in the
well-fed experimental group was 38.7%, although the latter might be an over-
estimation, because there was no sign of leveling off at this level.

Discussion

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The tidal effect of the water changes in Vagsb0pollen is restricted through the
narrow channel between Vagsb0pollen and the nearby Kviturdvikpollen, thereby
probably making the internal production of zooplankton the main source of food
for A.aurita. The biomass of zooplankton such as copepods was very low through-
out the investigation, so A.aurita was apparently under food-limited conditions,
although zooplankton not retained by the 180 um meshed net might have been
available. However, the most characteristic feature of the population was the high
abundance, small average size and size at maturity, all indicating food scarcity. A
survey of the literature data shows a large variability in population characteristics
(Yasuda, 1971; Hamner and Jenssen, 1974; Moller, 1980; Hernroth and Grondahl,
1985; van der Veer and Oorthuysen, 1985; Schneider, 1989; Lucas and Williams,
1994; Olesen et al., 1994; Schneider and Behrends, 1994; Ishii et al., 1995; Omori
et al., 1995; Lucas, 1996). We have compiled such data, especially related to how
open the environments occur, i.e. open, semi-enclosed and enclosed areas (Table
II). The locality and the pattern of population dynamics of A.aurita in this study
were similar to those from Kertinge Nor in Denmark (Olesen et al., 1994) and the
enclosed, man-made Horsea Lake in England (Lucas, 1996). The abundance of
A.aurita in Vagsb0pollen was considerably higher than that in most open-water
areas, but lower than that in Kertinge Nor and Horsea Lake (Table II). The semi-
enclosed and enclosed environments diverge from the open environments by
showing a range in abundance of 9-300 medusae m~3 compared to 0.01-1.5
medusae m~3 in the open environments. Accordingly, peak bell diameter reported
from open waters showed a range from 20 to 30 cm, whereas medusae from semi-
enclosed/enclosed areas reached a peak diameter of 4-10 cm, and the maximum
observed diameter showed the same trend. The few data available on minimum
size at maturity (Table II) do not contradict the hypothesis of smaller size in non-
open environments.
Schneider and Behrends (1994) suggested that the years of small-sized A.aurita
of high abundance and the years of large-sized individuals of low abundance
indicate the regulation mechanism of their density by food availability. In
Vagsb0pollen, food was apparently restricted to small hydromedusae and poten-
tially also to microzooplankton, although prey in the size range of ciliates and
rotifers do not seem to contribute substantially to the energy budget of A.aurita
(Bamstedt, 1990). The situation is thus similar to other non-open areas, where
food limitation is suggested as the explanation for the poor growth of medusae
(Olesen et al., 1994; Lucas, 1996). Through our experimental approach, we have
proved for the first time that food availability is really a controlling factor for the
813
oo

Table II. Aurelia aurita: review of the population studies

Area Location Abundance Max bell Max mean bell Min bell diameter References
index (ind. nr 3 ) diameter (cm) diameter (cm) at maturity (cm)
o Tokyo Bay, Japan nd nd 23 nd Ishii etal. (1995)
o Tokyo Bay, Japan 1.53 31 nd 8.5 Omori etal. (1995)
o Kiel Bight, western Baltic 0.16 30-33 22.5 nd Schneider and Behrends (1994)"
o Kiel Bight, western Baltic 0.006 39-42 30.3 nd Schneider and Behrends (1994) b
o Kiel Bight, western Baltic 0.23 nd 22 nd Schneider (1989)'
o Kiel Bight, western Baltic 0.04 nd 30 nd Schneider (1989)d
o Wadden Sea, Netherlands 0.25 nd 24 nd van der Veer and Oorthuysen (1985)'
o Wadden Sea, Netherlands 0.49 nd 20 nd van der Veer and Oorthuysen (1985)'
o Kiel Bight, western Baltic 0.33 44 19.7 nd Mdller (1980)
o Urazoko Bay, Japan Sea nd 31 22 7 Yasuda(1971)
s Southampton Water, England 8.71 15 10 6.4 Lucas and Williams (1994)
s Kertinge Nor, Denmark 300 nd 3.7 nd Olesen etat. (1994)
s Vagsb0pollen, Norway 22.3 11.8 8 5 Present study
e Horsea Lake, England 24.9 10.5 5.38 1.9 Lucas (1996)
Area index: o, open water; s, semi-enclosed water; e, enclosed water; nd, no data.
•Data of 1982.
b
Data of 1984.
c
Data of 1982.
d
Data of 1983 and 1984.
'Data of 1981.
'Data of 1982.

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 Food regulation ot Aurelia

growth of medusae in natural environments. The experiments also proved that

the maturation process is strongly governed by food availability, and the experi-
mental observations were thus in line with the field observations reported in the
literature (see Table II), where open waters show larger size at maturity than
semi-enclosed and enclosed areas. The same observations have been made from
Ninegret Pond, Charlestown, Rhode Island, where growth and size at maturity
are considerably lower in the semi-enclosed saltwater pond than outside in the
Block Island Sound (B.Sullivan, personal communication). Thus, food avail-

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ability seems to give a general explanation for both individual growth of the
medusa and its maturation, but it cannot explain why semi-enclosed/enclosed
areas have a high population abundance compared to open areas. Some possible
factors are discussed below.
First, planulae in open waters will be more dispersed, due to the larger area
and the higher degree of mixing. Secondly, part of the larval population in open
waters may settle in localities of unsuitable depth, thus being lost. Thirdly, a high
abundance of medusae in environments with restricted water exchange will also
result in a high abundance of planula larvae, and since the larval phase is short
(-7 days) it will lead to a high abundance of polyps that will produce a high abun-
dance of ephyrae in the next year. Finally, the other common scyphomedusan in
Scandinavian waters, Cyanea capillata, is known to be a main predator on
A.aurita (Bamstedt et al., 1994; Hansson, 1997), and it is not usually found in the
non-open environments that are commonly brackish, thereby reducing the
mortality of A.aurita.
Aurelia aurita thus seems to be a rather flexible species, being able to utilize
suitable food conditions for growth and reproduction, and easily adapt to
environments with reduced food sources through a reduction in somatic growth,
and an allocation of food energy to reproduction. This plasticity in food-energy
allocation makes it a strong competitor in diverse environments and explains its
worldwide success in coastal waters.

Acknowledgements
The report forms part of a research project financially sponsored by the Nor-
wegian Research Council. We sincerely thank the Ministry of Education, Science,
Sports, and Culture, Government of Japan, for a research scholarship to one of
us (H.I.) that made this study possible. We also thank Monica Bente Martinussen,
University of Bergen, and the staff at the marine biological field station for their
help.

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Fish., 37, 364-370.

Received on August 26, 1997; accepted on December 1, 1997

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